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245px, A flock of Dunlins_and_Red_knots.html" ;"title="Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders are birds of the order Charadriiformes commonly found along shorelines and mudflats that wikt:wade#Etymology 1, wade in order to foraging, forage for food (such as
insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to be in ...

insect
s or crustaceans) in the mud or sand. The term "wader" is used in Europe, while "shorebird" is used in North America, where "wader" may be used instead to refer to long-legged wading birds such as
stork Storks are large, long-legged, long-necked wading bird FIle:Vadare - Ystad-2021.jpg, 245px, A flock of Dunlins and Red knots Waders are birds of the order Charadriiformes commonly found along shorelines and mudflats that wikt:wade#Etymology ...

stork
s and
heron The herons are long-legged, long-necked, freshwater and coastal bird Birds are a group of warm-blooded vertebrates constituting the class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a ...

heron
s. There are about 210
species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individu ...

species
of wader, most of which live in wetland or coastal environments. Many species of Arctic and temperate regions are strongly migratory, but
tropical The tropics are the region of Earth surrounding the Equator. They are delimited in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S; these latitudes correspond to ...

tropical
birds are often resident, or move only in response to rainfall patterns. Some of the Arctic species, such as the
little stint The little stint (''Calidris minuta'') (or ''Erolia minuta''), is a very small wader Waders or shorebirds are bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathe ...
, are amongst the longest distance migrants, spending the non-
breeding season Seasonal breeders are animal species that successfully mating, mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, a ...
in the
southern hemisphere The Southern Hemisphere is the half (hemisphere Hemisphere may refer to: * A half of a sphere As half of the Earth * A hemispheres of Earth, hemisphere of Earth ** Northern Hemisphere ** Southern Hemisphere ** Eastern Hemisphere ** Western He ...

southern hemisphere
. Many of the smaller species found in
coastal The coast, also known as the coastline or seashore, is defined as the area where land meets the ocean The ocean (also the sea The sea, connected as the world ocean or simply the ocean The ocean (also the sea or the world ...

coastal
habitats, particularly but not exclusively the calidrids, are often named as "sandpipers", but this term does not have a strict meaning, since the
upland sandpiper The upland sandpiper (''Bartramia longicauda'') is a large sandpiper, closely related to the curlew The curlews (), genus ''Numenius'', are a group of nine species of birds, characterised by long, slender, downcurved bills and mottled brown p ...
is a grassland species. The smallest member of this group is the least sandpiper, small adults of which can weigh as little as and measure just over . The largest species is believed to be the
Far Eastern curlew The Far Eastern curlew (''Numenius madagascariensis'') is a large shorebird Waders or shorebirds are bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothl ...
, at about and , although the
beach thick-knee The beach stone-curlew (''Esacus magnirostris'') also known as beach thick-knee is a large, ground-dwelling bird that occurs in Australasia Australasia is a region In geography Geography (from Greek: , ''geographia'', literally "ear ...
is the heaviest at about .


Taxonomy

In the Sibley-Ahlquist taxonomy, waders and many other groups are subsumed into a greatly enlarged order
Ciconiiformes Storks are large, long-legged, long-necked wading bird Waders or shorebirds are bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless beaked jaws, the ...
. However, the classification of the Charadriiformes is one of the weakest points of the Sibley-Ahlquist taxonomy, as
DNA–DNA hybridization DNA–DNA hybridization generally refers to a molecular biology technique that measures the degree of genetic similarity between pools of DNA sequences. It is usually used to determine the genetic distance between two organisms. This has been used ...
has turned out to be incapable of properly resolving the interrelationships of the group. Formerly, the waders were united in a single suborder ''Charadrii'', but this has turned out to be a "
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical s ...
", uniting no fewer than four charadriiform lineages in a
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The group is said to be paraphyleti ...

paraphyletic
assemblage. However, it indicated that the plains wanderer actually belonged into one of them. Following recent studies (Ericson et al., 2003; Paton et al., 2003; Thomas et al., 2004a, b; van Tuinen et al., 2004; Paton & Baker, 2006), the waders may be more accurately subdivided. The waders are a group of two Charadriiformes
suborder In biological classification, the order ( la, wikt:ordo#Latin, ordo) is # a taxonomic rank used in the classification of organisms and recognized by the nomenclature codes. The well-known ranks in descending order are: life, domain (biology), do ...
s which include 13
families In human society, family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of families is to maintain the w ...
. Species in the third Charadriiforme suborder,
Lari The suborder Lari is the part of the order Charadriiformes Charadriiformes (, from '' Charadrius'', the type genus Type may refer to: Science and technology Computing * Typing Typing is the process of writing or inputting text by pressing ...

Lari
, are not considered as waders. * Suborder Charadrii ** Family
Burhinidae The stone-curlews, also known as dikkops or thick-knees, consist of nine species within the family Burhinidae, and are found throughout the tropical and temperate parts of the world, with two or more species occurring in some areas of Africa, Asia ...
– stone-curlews, thick-knees (10 species) ** Family Pluvianellidae – Plains-wanderer ** Family Chionidae – sheathbills (2 species) ** Family Pluvianidae – Egyptian plover ** Family
Charadriidae The bird family Charadriidae includes the plover Plovers ( or ) are a widely distributed group of wader, wading birds belonging to the subfamily Charadriinae. Description There are about 66 species in the subfamily, most of them called "pl ...
– plovers (68 species) ** Family
Recurvirostridae The Recurvirostridae are a family of bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, ...
– stilts, avocets (10 species) ** Family Ibidorhynchidae – ibisbill ** Family Haematopodidae – oystercatchers (12 species) * Suborder Scolopaci ** Family Rostratulidae – painted-snipes (3 species) ** Family
Jacanidae The jacanas (sometimes referred to as Jesus birds or lily trotters) are a group of tropical waders in the family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or ...
– jacanas (8 species) ** Family
Pedionomidae The plains-wanderer (''Pedionomus torquatus'') is a bird, the Monotypic taxon, only representative of family Pedionomidae and genus ''Pedionomus''. It is endemic to Australia. The majority of the remaining population is found in the Riverina regi ...
– plains-wanderer ** Family Thinocoridae – seedsnipes (4 species) ** Family
Scolopacidae Sandpipers are a large family, Scolopacidae, of wader 245px, A flock of Dunlins and Red knots Waders are birds of the order Charadriiformes commonly found along shorelines and mudflats that wikt:wade#Etymology 1, wade in order to foraging, ...
– sandpipers, snipes (98 species)


Characteristics

Shorebirds is a blanket term used to refer to multiple bird species that live in wet, coastal environments. Because most these species spend much of their time near bodies of water, many have long legs suitable for wading (hence the name ‘Waders’). Some species prefer locations with rocks or mud. Many shorebirds display migratory patterns and often migrate before breeding season. These behaviors explain the long wing lengths observed in species, and can also account for the efficient metabolisms that give the birds energy during long
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
s."Explore the World With Shorebirds." U.S. Fish and Wildlife Service, 1 Aug. 2004. Web.<http://www.fws.gov/alaska/external/education/pdf/Chap4.pdf>. The majority of species eat small
invertebrate Invertebrates are animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This includes all animals apart from the chordata, chordate subphylum vertebrate, Vertebra ...
s picked out of
mud Mud is soil Surface-water- gley developed in glacial till, Northern Ireland.">Northern_Ireland.html" ;"title="glacial till, Northern Ireland">glacial till, Northern Ireland. Soil is a mixture of organic matter, minerals, gases, liquids, a ...

mud
or exposed soil. Different lengths of bills enable different species to feed in the same habitat, particularly on the coast, without direct competition for food. Many waders have sensitive nerve endings at the end of their bills which enable them to detect prey items hidden in mud or soft soil. Some larger species, particularly those adapted to drier habitats will take larger prey including
insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to be in ...

insect
s and small
reptiles Reptiles, as most commonly defined, are the animals in the class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or ...

reptiles
.


Sexual dimorphism

Shorebirds, like many other animals, exhibit
phenotypic In genetics Genetics is a branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular inter ...

phenotypic
differences between males and females, also known as
sexual dimorphism Sexual dimorphism is the condition where the sex Sex is either of two divisions, typically male Male (♂) is the sex of an organism that produces the gamete known as sperm. A male gamete can fuse with a larger female gamete, or ovum, in ...
. In shorebirds, various sexual dimorphisms are seen, including, but not limited to, size (e.g. body size, bill size), color, and agility. In
polygynous Polygyny (; from Neoclassical Greek πολυγυνία from πολύ- ''poly-'' "many", and γυνή ''gyne'' "woman" or "wife") is the most common and accepted form of polygamy Polygamy (from Greek language, Late Greek , ''polygamí ...
species, where one male individual mates with multiple female partners over his lifetime, dimorphisms tend to be more diverse. In
monogamous Monogamy ( ) is a form of dyadic relationship Relationship most often refers to: * Interpersonal relationship The concept of interpersonal relationship involves social associations, connections, or affiliations between two or more peo ...
species, where male individuals mate with a single female partner, males typically do not have distinctive dimorphic characteristics such as colored feathers, but they still tend to be larger in size compared to females. The suborder Charadrii displays the widest range of sexual dimorphisms seen in the order Charadriiformes.Székely, Tamás, John D. Reynolds, and Jordi Figuerola. 2000. Sexual Size Dimorphism In Shorebirds, Gulls, And Alcids: The Influence Of Sexual And Natural Selection. 54(4): 1404-413

However, cases of sexual monomorphism, where there are no distinguishing physical features besides external genitalia, are also seen in this order.Lindenfors, P., T. Szekely, and J. D. Reynolds. "Directional Changes in Sexual Size Dimorphism in Shorebirds, Gulls and Alcids." Journal of Evolutionary Biology J. Evolution Biol: 930-38. Print.


Sexual selection

One of the biggest factors that leads to the development of sexual dimorphism in shorebirds is
sexual selection Sexual selection is a mode of natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrat ...
.Szekely, T., R. P. Freckleton, and J. D. Reynolds. "Sexual Selection Explains Rensch's Rule of Size Dimorphism in Shorebirds." ''Proceedings of the National Academy of Sciences'' (2004): 12224-2227. Print. Males with ideal characteristics favored by females are more likely to reproduce and pass on their genetic information to their offspring better than the males who lack such characteristics. Mentioned earlier, male shorebirds are typically larger in size compared to their female counterparts. Competition between males tends to lead to sexual selection toward larger males and as a result, an increase in dimorphism. Bigger males tend to have greater access (and appeal) to female mates because their larger size aids them in defeating other competitors. Likewise, if the species exhibits gender role reversal (where males take on roles traditionally done by females such as childcare and feeding), then males will select female mates based on traits that are the most appealing. In the Jacana species, females compete with each other for access to male mates, so females are larger in size. Males choose female mates based on who presents herself as the strongest and who 'owns' the most territory.


Natural selection

Another factor that leads to the development of dimorphisms in species is
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
. Natural selection focuses on traits and the environment's response to the traits in question; if the said trait increases the overall fitness of the individual possessing it, then it will be 'selected' and eventually become a permanent part of the population's gene pool. For example, depending on the food available in a shorebird specie's respective
niche Niche may refer to: Science *Developmental niche{{third-party, date=October 2020 The developmental niche is a theoretical framework for understanding and analyzing how culture Culture () is an umbrella term which encompasses the social behav ...

niche
, bigger bill sizes may be favored in all individuals. This would essentially lead to monomorphism within the species but is subject to change once sexual selection acts on the trait. Sexual selection could give rise to males with relatively larger bills than females if males used their bills to compete with other males. If larger bill size assisted the male in gathering resources, it would also make him more attractive to female mates.


See also

*
Hybridisation in shorebirdsHybrid (biology), Hybridisation in wader, shorebirds has been proven on only a small number of occasions; however, many individual shorebirds have been recorded by birdwatchers worldwide that do not fit the characters of known species. Many of these ...
* List of Charadriiformes by population


References


Sources

* Ericson, P. G. P.; Envall, I.; Irestedt, M.; & Norman, J. A. (2003). Inter-familial relationships of the shorebirds (Aves: Charadriiformes) based on nuclear DNA sequence data. '' BMC Evol. Biol.'' 3: 16.
PDF fulltext
*Pandiyan, J. and S. Asokan. 2015. Habitat use of pattern of tidal mud and sandflats by shorebirds (charadriiformes) Wintering in southern India. Coastal Conservation https://doi.org/10.1007/s11852-015-0413-9. * Paton, Tara A.; & Baker, Allan J. (2006). Sequences from 14 mitochondrial genes provide a well-supported phylogeny of the Charadriiform birds congruent with the nuclear RAG-1 tree. ''Molecular Phylogenetics and Evolution'' 39(3): 657–667. (HTML abstract) * Paton, T. A.; Baker, A. J.; Groth, J. G.; & Barrowclough, G. F. (2003). RAG-1 sequences resolve phylogenetic relationships within charadriiform birds. ''Molecular Phylogenetics and Evolution'' 29: 268–278. (HTML abstract) * Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás (2004a). Phylogeny of shorebirds, gulls, and alcids (Aves: Charadrii) from the cytochrome-''b'' gene: parsimony, Bayesian inference, minimum evolution, and quartet puzzling. ''Molecular Phylogenetics and Evolution'' 30(3): 516–526. (HTML abstract) * Thomas, Gavin H.; Wills, Matthew A.; & Székely, Tamás (2004b). A supertree approach to shorebird phylogeny. '' BMC Evol. Biol.'' 4: 28.
PDF fulltextSupplementary Material
* van Tuinen, Marcel; Waterhouse, David; & Dyke, Gareth J. (2004). Avian molecular systematics on the rebound: a fresh look at modern shorebird phylogenetic relationships. ''Journal of Avian Biology'' 35(3): 191–194
PDF fulltext
* Explore the World With Shorebirds. (2004). U.S. Fish and Wildlife Service. Web. http://digitalmedia.fws.gov/cdm/ref/collection/document/id/1598 * Lindenfors, P.; Szekely, T.; and Reynolds, J. D. (2003). Directional Changes in Sexual Size Dimorphism in Shorebirds, Gulls and Alcids. Journal of Evolutionary Biology J Evolution Biol: 930–38. Print. * Szekely, T.; Freckleton, R.; & Reynolds, J. (2004). Sexual selection explains Rensch's rule of size dimorphism in shorebirds. ''Proceedings of the National Academy of Sciences.'' 101(33): 12224–12227. * Szekely, Tamas; John D. Reynolds; and Jordi Figuerola. (2000) Sexual Size Dimorphism in Shorebirds, Gulls, and Alcids: The Influence of Sexual and Natural Selection. Evolution 54(4): 1404–413.


External links

* {{Authority control * * * Bird common names